Summary 1. Crustaceans have a high content of calcium, which is chiefly located in the skeleton as calcium carbonate. Calcium is generally the most abundant cation in the body. 2. During intermoult, the exoskeleton is usually fully calcified and the animal is in calcium equilibrium with its environment. 3. In the premoult stages calcium is resorbed from the skeleton and may be lost to the environment or stored within the body. Typically, losses are high and storage is small in aquatic species, whilst most terrestrial forms store much larger amounts of calcium and losses are reduced. Loss of calcium in soluble form by aquatic species must be by outward transport across the gills. 4. Calcium is stored in a variety of different ways, usually with a common taxonomic theme. The main forms are as calcium phosphate granules in cells of the midgut gland (Brachyura), gastroliths (Astacidea and some Brachyura), the haemocoel (some Brachyura) the posterior midgut caeca (Amphipoda) and the ventral portion of the body generally in the Isopoda. 5. At ecdysis, the skeleton is shed and the calcium remaining in it is lost from the body. 6. Recalcification begins immediately, or shortly after, ecdysis using calcium mobilized from the stores. Simultaneously, or when the stores are exhausted, other sources of calcium are utilized. These are calcium in the water (aquatic species), the food (aquatic and terrestrial species) and the exuviae (chiefly terrestrial species). 7. Marine species store little calcium and must obtain the bulk of their requirement (ca. 95%) from the water. Fresh‐water species also store little calcium but have, seemingly, adapted to the lower availability of calcium by increasing the affinity of the calcium‐absorbing mechanism. The rates of uptake of calcium are consequently similar in marine and fresh‐water species. 8. A high degree of storage is essential for terrestrial crustaceans as they do not have access to a large aquatic reservoir of calcium. These large reserves enable the animals to reach an advanced stage of calcification, allowing the resumption of foraging and feeding necessary for completion of calcification. 9. The control of calcium metabolism during the intermoult cycle is poorly under stood. β Ecdysone appears to control the resorption of calcium and the formation of calcium stores during premoult, but the mechanism of control of calcium metabolism during postmoult and intermoult is unknown. 10. The concentration of calcium in the haemolymph of most species is high, but a large proportion of this is in non‐ionized form. In premoult, total calcium levels rise as a result of calcium resorption but little change occurs in the concentration of ionized calcium. Postmoult generally sees a fall in blood calcium, sometimes below the intermoult value.
This paper reviews the nutritional ecology, the digestive physiology, and biochemistry of herbivorous land crabs and the adaptations that they possess towards a diet of plant material. Land crab species that breathe air and forage out of water can be divided into three feeding specialisations: primarily carnivorous, deposit feeders feeding on micro-organisms and organic matter in the sediment, and herbivores consuming mainly plant material and its detritus. The last forms the focus of this review. The diets of the herbivores are low in nitrogen and high in carbon, are difficult to digest since they contain cellulose and hemicellulose, and may disrupt digestion due to the presence of tannins. Herbivorous crustaceans are able to efficiently utilise plant material as their primary nutrient source and are indeed able to meet their nitrogen requirements from it. Herbivorous land crabs display a range of adaptations towards a low nitrogen intake and these are discussed in this review. They also appear to endogenously produce cellulase and hemicellulase enzymes for the digestion of cellulose and hemicellulose. Generalised and specific adaptations allow them to inhibit the potentially negative digestive effects of tannins. To digest plant material, they possess a plastic digestive strategy of high food intake, short retention time, high assimilation of cell contents, and substantial digestion of cellulose and hemicellulose.
The robber crab (Birgus latro), also known as the coconut crab, is the world's largest land-living arthropod, with a weight reaching 4 kg and a length of over half a meter. Apart from the marine larval stage, this crab is fully terrestrial, and will actually drown if submerged in water. A transition from sea to land raises dramatically new demands on the sensory equipment of an animal. In olfaction, the stimulus changes from hydrophilic molecules in aqueous solution to mainly hydrophobic in the gaseous phase. The olfactory system of land crabs thus represents an excellent opportunity for investigating the effects of the transition from sea to land. Have land crabs come to the same solutions as other terrestrial animals, or is their olfactory sense characterized by unique innovations? Here, we show that the robber crab has evolved an olfactory sense with a high degree of resemblance to the insect system. The similarities extend to physiological, behavioral, and morphological characters. The insect nose of the robber crab is a striking example of convergent evolution and nicely illustrates how similar selection pressures result in similar adaptation.
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